A Revision of Solanum Section Herpystichum-Solanaceae

8/14/2019 A Revision of Solanum Section Herpystichum-Solanaceae

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A Revision of SolanumSection Herpystichum

Author(s): Eric J. Tepe and Lynn Bohs

Source: Systematic Botany, 36(4):1068-1087. 2011.

Published By: The American Society of Plant Taxonomists

URL: http://www.bioone.org/doi/full/10.1600/036364411X605074

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Systematic Botany (2011), 36(4): pp. 10681087 Copyright 2011 by the American Society of Plant TaxonomistsDOI 10.1600/036364411X605074

1068

The genus Solanum L. is found worldwide in temperate

and tropical habitats and, with ca. 1,500 species, is one ofabout a dozen giant genera with over 1,000 species (Frodin2004). The informally named Potato clade is one of thelargest of the 1113 well-supported clades within Solanum,as revealed by molecular data (Bohs 2005; Weese and Bohs2007). Because it contains the cultivated species potato(S. tuberosumL.), tomato (S. lycopersicumL.), and the less well-known crop pepino or pepino dulce (S. muricatumAiton), thePotato clade is also one of the most intensively studied cladesofSolanum. Despite this focus on the crop species and theirclose relatives, other members of the clade have received lit-tle taxonomic attention. In fact, some of the less well-knowngroups that make up the Potato clade, such as Solanumsec-tionsHerpystichumBitter and PteroideaDunal, have only been

recently recognized as belonging there based on morphology(Child 1990) and molecular data (Bohs 2005; Weese and Bohs2007). Knapp and Helgason (1997) provided a taxonomicrevision of sect. Pteroidea, but sect.Herpystichumhas not beenstudied taxonomically. The present study is a revision ofSolanumsect.Herpystichum.

Solanum sect. Herpystichum contains 10 species restrictedto wet habitats, ranging from southern Mexico to northernPeru. All species of the section are ground-trailing or climbingvines that root at the nodes; pubescence, when present, is ofunbranched hairs. Most species have fruits that are flattenedperpendicular to the septum to varying degrees. Many spe-cies of the section have globose, apically pointed flower buds,distinctive within Solanum. In general, sect.Herpystichumhas

plurifoliate sympodial units and extra-axillary inflorescences.Many species of sect. Herpystichumare narrowly distributed,relatively inconspicuous, and tend to be rare in the habitatswhere they occur. Consequently, they are among the least col-lected species ofSolanum.

Two distinct groups are apparent within sect.Herpystichum.One, the ground-trailing species, have herbaceous stemsthat creep along the ground and have either simple or com-pound leaves with long petioles (>3 cm). This group includesS. dalibardiforme, S. limoncochaense, S. pentaphyllum, S. phase-oloides, and S. trifolium(Figs. 1A -E, 2 A-B). The other groupcomprises climbing species with herbaceous to woodystems, all with simple leaves with short petioles (


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2011] TEPE AND BOHS: SOLANUM SECT. HERPYSTICHUM 1069

Fig. 1. Flowers fruits, and habit of Solanumsect.Herpystichum: A. Solanum limoncochaense, habit, flower, and bud. B. S. limoncochaensefruit in situ andC. in cross section (E. J. Tepe & S. Stern 2627, QCNE). D. S. trifoliumflower and leaf (E. J. Tepe & S. Stern 2682, QCNE). E. S. phaseoloideshabit, flower, andleaf (photo by W. H. Haber). F. S. crassinerviumhabit (E. J. Tepe & S. Stern 2729 , QCNE), and G. flowers (S. Stern & E. J. Tepe 400, QCNE). H. S. evolvulifo-liumhabit and flower, and I. fruit (immature) (E. J. Tepe & S. Stern 2671 , QCNE). J. S. pacificuminflorescence with flowers, buds, and fruit (immature), andK. habit (E. J. Tepe et al. 2696, QCNE). L. S. loxophyllumhabit, note adventitious roots at nodes, and M. older stem with inflorescences, note figure eight-shaped stem (E. J. Tepe & S. Stern 2726, QCNE). Inflorescences emerge from the cleft between the two lobes of the stem.


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1070 SYSTEMATIC BOTANY [Volume 36

numbers are cited with the herbarium acronym followed by the sheetnumber (i.e. MO1781232), or in the case of QPLS, their generic abbrevia-tion followed by the sheet number (i.e. QPLSSOLA0192); barcodes arepreceded by the herbarium acronym (i.e. GG00104280, or AGH00077619

since all herbaria incorporated into the Harvard University Herbariaare cited separately but given GH barcodes). Some specimens have

both sheet numbers and barcodes and in these cases, only the barcodeis listed.

Fig. 2. Flowers, fruits, and habit of Solanumsect.Herpystichum. A. Solanum pentaphyllum, habit and flower (from O. Haught 2551, F). B. S. dalibardiforme,habit and flower (from K. von Sneidern 3121, US). C. S. dolichorhachis, habit, flower, and immature fruit ( from F. H. Eggers 14641, A).


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We have followed the morphological species concept or morphologi-cal cluster species concept (Mallet 1995) in delimiting the species recog-nized in this study. Individuals that clustered together based on a commonset of morphological characters and that were separated from other clus-ters by gaps in the morphological continuum were considered species.In most cases, species that we have identified based on morphology alsooccupy coherent geographic ranges. Our goal was to produce a classifica-tion that both reflects the biology of the species, and that can be used byspecialists and non-specialists alike to identify the taxa in the group.

The phylogeny presented here is summarized from Tepe et al. (2011)

and includes DNA extracted from fresh, silica gel-dried leaf material,or herbarium specimens for 25 accessions of Solanum. These accessionsincluded 19 from sect.Herpystichumand six were outgroups from relatedsections. Analyses included data from seven nuclear (ITS, GBSSI, and fiveCOSII) and three plastid (psbAtrnH, trnTtrnF, and trnStrnG) regions.Details of DNA extraction, primers, and PCR conditions are presented inTepe et al. (2011). Phylogenetic relationships in the tree presented wereestimated using Bayesian inference (Huelsenbeck and Ronquist 2001;Ronquist and Huelsenbeck 2003) of a partitioned, concatenated dataset.Substitution model parameters were determined with MrModeltest 2.2(Nylander 2004). Further details of analyses can be found in Tepe et al.(2011).

Results

Taxonomic HistoryMichel Flix Dunal described the first

species of this group, the Ecuadorean S. trifolium, in 1852,based on a collection from Ruiz and Pavns exploration ofChile and Per s. l. (Tafalla and Estrella 1989). He allied thespecies to the compound-leaved members of what we nowknow as Solanumsect. Pteroidea, under his Polybotryongroup(Dunal 1852). The second species described was S. phase-oloides, by Hellmuth Polakowsky in 1877 from his own CostaRican collection. Polakowsky followed Dunals classifica-tion scheme, placing his new species in Polybotryon, and rec-ognized the close relationship between his new species andS. trifolium. John Donnell Smith described S. olivaeforme(a syn-onym ofS. phaseoloides) in 1889, but suggested a relationshipwith S. tripartitumDunal, a member of the Morelloid clade(Weese and Bohs 2007), based on the fruits among other dif-

ferences. Solanum evolvulifolium was the first of the climb-ing species to be described, by Jesse Greenman in 1904, basedon a Costa Rican collection, but without commentary on itsrelationships (Donnell Smith 1904).

Most of the remaining species were described by GeorgBitter (1912a, 1913a, b) in the early 20th century. In his descrip-tion of S. loxophyllum, Bitter (1912a) likened it in habit andmorphology to members of sects. Anarrhichomenum Bitterand Polybotryon (G. Don) Bitter (= sect. Pteroidea Dunal proparte; see Knapp and Helgason 1997 for more details), butsuggested that it may be distinct enough to warrant a newsection of its own: sect. Loxophylla Bitter (ined.). Later thesame year he explicitly excluded S. loxophyllum from sect.Polybotryon (Bitter 1912b), but he did not comment further

on its relationships because the species was still insufficientlyknown. Solanum dalibardiformeand S. dolichorhachiswere pub-lished simultaneously in 1913 (Bitter 1913a). Bitter mentioneda possible link between S. dolichorhachis and S. loxophyllum,

but emphasized that he had not yet seen the flowers ofS. lox-ophyllumnor the fruits of S. dolichorhachis. He also suggestedan association among S. dalibardiforme, S. phaseoloides, S. trifo-lium, and S. ionidiumBitter. Solanum ionidiumis a member ofsect. Anarrhichomenumand is distinct from the other specieslisted here; however, the type is a rather small fragment of anherbaceous stem tip, which is not representative of the actualhabit of this typically woody species. Finally, later in 1913Bitter published S. pentaphyllum, suggesting a close relation-

ship with S. phaseoloides(Bitter 1913b). Bitter (1921) mentionedthe sectional nameHerpystichumin reference to S. pentaphyl-lum, S. phaseoloides, and S. trifolium, which he differentiatedfrom sect. Polybotryonbased on the extra-axillary position ofthe inflorescences of the former. In this passage, Bitter citessect.Herpystichumas if it had been published previously, butwe have not been able to locate any previous mention of thesection. This apparent reference appears to have caused some

confusion, and some authors have cited several dates for thepublication of sect.Herpystichum(see Seithe 1962). Althoughnot a formal sectional description, the information includedin the 1921 passage, however, satisfies the retroactive ICBNrules for valid publication of names (McNeill et al. 2006) and,thus, we consider SolanumsectionHerpystichumto have beenvalidly published by Bitter in 1921.

Seithe (1962) grouped the ground-trailing species S. penta-phyllum, S. phaseoloides, and S. trifoliumin her circumscriptionof sect. Herpystichum, but excluded the climbing species S.dolichorhachisand S. evolvulifoliumto unspecified status withinher subg. Solanum. Likewise, DArcy (1973) placed the ground-trailing species S. phaseoloideswithin sect. Herpystichum, butplaced S. evolvulifolium in sect. Anarrhichomenum. Similarly,

Child (1990), in his synopsis of DArcys subg. Potatoe (G.Don) DArcy, grouped the ground-trailing species in sect.Herpystichum, including, as did Bitter, S. ionidium, and follow-ing DArcy, placed S. evolvulifoliumin sect.Anarrhichomenum.Nevertheless, Child was the first to ally Herpystichumto thepotatoes s. l. He did not provide explicit reasons for the rela-tionship, but saw well-developed compound leaves, whichare uncommon in Solanum, as a uniting feature of subg.Potatoe(Child 1990). Nee (1999) grouped the ground-trailingand climbing species for the first time under his circumscrip-tion of sect.Herpystichum, but did not state his reasons for thegrouping.

The remaining three species were described by Tepe (Tepeand Bohs 2009). Many species of sect.Herpystichumhave nar-

row distributions and are rare, thus we think it is likely thatadditional species exist in poorly collected areas, especiallysouthern Ecuador and northern Peru.

Morphology and Natural HistoryHabitMembers ofsect. Herpystichum are node-rooting vines. Five species trailalong the ground on the forest floor or in clearings, often indense patches, but are also weak climbers that can scrambleover fallen trees. Solanum crassinervium, S. evolvulifolium, andS. pacificumare stronger climbers, frequently climbing under-story trees, and S. loxophyllum,and possibly S. dolichorhachis,can be strong climbers. Most species are entirely herbaceous(S. dalibardiforme, S. limoncochaense, S. pacificum, S. pentaphyl-lum, S. phaseoloides, and S. trifolium); however, stems of someof the climbing species are strongly woody (S. dolichorhachis,S. evolvulifolium, and loxophyllum), but with little accumulatedsecondary growth. Stems of S. crassinervium are woody, butalso somewhat fleshy, and are the thickest stems in the sec-tion. Older stems ofS. loxophyllumare shaped like a numeral8 in cross-section (referred to herein as 8-shaped; Fig. 1M).Otherwise, the stems, herbaceous and woody, are terete andslender. Sympodial units are plurifoliate in most species, withconsiderable vegetative growth between inflorescences. Theexception to this rule is S. crassinerviumin which the sympo-dial units are often unifoliate.

TrichomesTrichomes are unbranched, uniseriate, mul-ticellular, and non-glandular. Three species, S. pentaphyllum,S. phaseoloides, and S. trifolium have distinctive, markedly


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tapered, uniseriate trichomes on the upper surfaces of theleaves that are 45 times wider at the base than the trichomeson the stems, petioles and lower leaf surfaces of the sameindividuals. Trichomes on other parts of those species,including the leaf veins adaxially, and on the remaining spe-cies, are slender and similar to the non-glandular, multicellu-lar trichomes of many other Solanumspecies (e.g. Seithe andAnderson 1982). The degree of pubescence is variable among

collections ofS. evolvulifolium,which range from nearly gla-brous to densely pubescent, but is relatively constant in otherspecies. Pubescence of short hairs (mostly 3 cm), whereas thoseof the climbing species are much shorter (nearly sessile to1.5 cm). The leaf blades, veins, and petioles of all species aremore or less punctate with deposits of crystal sand, whichare visible as small whitish dots (referred to herein as sand-punctate). These idioblasts filled with crystal sand are foundin several groups within Solanumas well as other groups ofangiosperms (Metcalfe and Chalk 1950; Whalen et al. 1986;Bohs 1990; Knapp 1992; Aliyu Aliero et al. 2006). Section

Herpystichumlacks the pseudostipules and interjected leaflets

(minute leaflets scattered among the larger leaflets, often seenon cultivated potato and tomato plants) that are present onmany groups within the Potato clade (Correll 1962).

InflorescencesInflorescences, like those throughoutSolanum, are morphologically terminal and the associated axil-lary bud continues the growth of the main stem axis (Danert1958). In all species, most inflorescences are extra-axillary, butthey can also be close enough to the node that they appearaxillary or leaf-opposed. Because inflorescences in Solanumare terminal, the variable position with respect to the leafaxil is likely the result of concaulescence of stem, petiole, andinflorescence tissues as is the case in sect. Pteroidea (Danert1967; Child 1979). Adventitious roots are frequently associ-ated with the attachment point of the extra-axillary inflores-

cences in the ground-trailing species. The inflorescences areslender and unbranched in most species, but occasionallyonce or twice branched in S. crassinervium. Inflorescencesin the majority of species are few-flowered (i.e. 210 flow-ers), but can have up to 50 flowers in S. evolvulifolium andS. pacificum, and > 100 in S. dolichorhachis. Only one or twoflowers are open at one time on an inflorescence, but thetotal number of flowers per inflorescence can be counted

by the scars left behind once the old flowers or fruits havefallen. Because the inflorescences of S. crassinervium can be

branched, they typically have more simultaneously openflowers. The pedicels of all species are articulated at the

base.

FlowersThe flower buds of sect. Herpystichum have adistinctive shape within Solanum. They are globose with anacuminately pointed apex resulting in a somewhat onion-shaped bud (see Fig. 1A). This shape is manifest in later stagesof bud development, and we have seen it in all but two species(S. dalibardiformeand S. trifolium). These two species, however,are represented by few collections, and the available buds areall at early stages. Thus, it is possible that onion-shaped buds

are characteristic of all species of sect.Herpystichum.The flowers are pentamerous and apparently perfect inall species. They are actinomorphic, with the exception ofS. crassinerviumin which the style is deflected to one side insome flowers. The anthers are yellow and equal in length,or only slightly unequal. The filaments are glabrous and notunited. The corollas are stellate in eight species (Figs. 1A, E,G, H, J, M), but there is a moderate to considerable amountof interpetalar tissue in S. dalibardiforme (Fig. 2B) and S. tri-

folium (Fig. 1D) resulting in rotate-stellate to pentagonal torotate corollas. Petals of all species are ciliate, and are rarelypubescent abaxially. Flower color in most species ranges fromgreenish-white to white to creamy-white, but the flowers ofS. trifoliumare pure blue-violet. Flower color in S. evolvulifo-

liumis variable among individuals and ranges from white toviolet, with frequent mottling. Flower color may also be vari-able in S. dalibardiformeand S. dolichorhachis, but we have notseen these two species in the field and they are known fromfew specimens.

FruitsFruits are variable among species of sect. Herpys-tichum, and range from more or less globose to stronglyflattened and arrowhead-shaped (Figs. 1B, C). All fruits, how-ever, have a tendency toward some degree of flattening andare always flattened perpendicularly to the septum. Fruitsflattened perpendicularly to the septum are rare in Solanumand this character, although not well developed in some spe-cies, helps to distinguish sect. Herpystichumfrom the rest ofthe genus.

In general, the fruits are glabrous, but those of S. limonco-chaensemay have some widely scattered hairs when young.Collections with mature fruits are rare, but green fruits prob-ably mature to orange or red in the climbing species, whereasthose of the ground-trailing species mature to yellow mottledwith brown (S. phaseoloides), bronze-brown (S. limoncochaense;Fig. 1B), or blackish-purple (S. pentaphyllum). Little is knownabout seed dispersal in species of sect.Herpystichum, but thefruits of S. limoncochaense, the only species with mature fruitsthat we have been able to observe in the field, are stronglyfragrant with a sweet, heavy scent. The color, scent, and posi-tion (i.e. lying on the ground) suggest that they are likely to bedispersed by agoutis or other terrestrial mammals (W. Haber,pers. comm.). The labels of two collections ofS. pentaphyllum

state that the fruits dig themselves into the ground as theymature; however, no other labels mention this phenomenonand we have not observed this species in the field. Additionalwork is necessary to determine whether S. pentaphyllum is,in fact, geocarpic. Stone cell aggregates, common in somegroups of Solanum (e.g. sect. Regmandra; Bennett 2008) areabsent from the fruits of sect.Herpystichum.

Habitat and Geographic DistributionSpecies of sect.Herpystichum are native to tropical America and are foundfrom southernmost Mexico (Chiapas), Belize, and Guatemalato northern Peru, from about 17N to 7S (Fig. 3). Speciesdiversity is highest between about 5N to 130S (see indi-vidual species descriptions). In South America, they occur in


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high- (mostly over 2,500 m) to mid-elevation (6502,500 m)Andean forests, or occur in the lowland forests to the east andwest of the Andes (sea level to mostly below 650 m). One spe-cies, S. pentaphyllum, extends into Venezuela along the Andes(Cordillera de Mrida) to the Cordillera de la Costa. With theexceptions ofS. evolvulifolium, S. pentaphyllum, and S. phase-oloides, species of sect. Herpystichum are narrow endemicsand rare (although some are locally abundant). Several spe-

cies appear to be sensitive to disturbance and are not found inhighly altered habitats, whereas others are frequently foundon forest edges, along roadsides and trails, or in pastures andother clearings. Although several species appear to thrive inopen, disturbed areas, species in this section are characteristi-cally understory plants of humid rainforests, premontane wetforests, or cloud forests. They are absent from dry or season-ally dry habitats.

Phylogenetic RelationshipsMolecular data supportsect. Herpystichum as a member of the Potato clade (Bohs2005; Weese and Bohs 2007). A more detailed phyloge-netic study of the Potato clade is currently underway (Tepeand Bohs unpubl. data) and preliminary analyses indi-cate five well-supported sub-clades that correspond to the

traditionally-recognized sections Anarrhichomenum, Basar-thrum,Herpystichum, and Pteroidea, and a clade containing sec-tions Etuberosum (Bukasov & Kameraz) A. Child, Juglandifolia(Rydberg) A. Child, Lycopersicoides (A. Child) Peralta and theeconomically important sections Lycopersicon (Miller) Wettsteinand PetotaDumort (see Spooner et al. 1993; Peralta et al. 2008and included references for details of relationships amongthese groups). Within the Potato clade, sectionsHerpystichumand Pteroideaare strongly supported as sister taxa (Fig. 4 ).

Tepe et al. (2011) provided a 10-marker phylogenetichypothesis for all 10 species of sect. Herpystichum and out-groups (Fig. 4). These results support two major lineageswithin sect.Herpystichum. One clade is composed of S. limon-

cochaense, S. pentaphyllum, S. phaseoloides, and S. trifolium.These are all ground-trailing species and share numerouscharacters, including markedly flattened fruits, white stellatecorollas in all species (except S. trifolium, which has rotate,

blue-violet flowers), and wide-diameter hairs on the adaxialleaf surfaces (except for S. limoncochaense, which is glabrous).Solanum dalibardiforme, the remaining ground-trailing spe-cies, is not included in this clade, but rather is sister to theother major lineage, which is composed of all of the simple-leaved climbing species. This placement is interesting becauseS. dalibardiformehas simple leaves like the climbing species,

but shares the ground-trailing habit and long petioles with

the ground-trailing species. Furthermore, its corolla is rotate-stellate to pentagonal to rotate, a character shared only withS. trifolium within the section. The node joining S. dalibar-diformeto the climbing species, however, has the lowest sup-port of any node in the tree.

The simple-leaved climbing species are a strongly-supported monophyletic group. Shared characters includea climbing habit, woody stems in all species except S. pacifi-cum, white to pink or violet stellate corollas, and a tendencytoward oblique leaf bases. All of these species have distinc-tive, short trichomes on the leaf midveins adaxially (exceptfor S. pacificum, which is completely glabrous). Our resultsindicate that S. crassinerviumand S. loxophyllumhave evolved

Fig. 3. Distribution of Solanumsect.Herpystichum.

Fig. 4. Summary phylogeny of Solanumsect. Herpystichumbased onbased on plastid psbA-trnH, trnT-trnF, and trnS-trnG, and nuclear waxy,ITS, cos1C, cos5, cos9B, cos10B, and cos11 sequences (E. J. Tepe et al.2011). For clarity, replicate accessions of all species except for S. evolvu-lifoliumand S. pentaphyllumhave been omitted. Provenance is indicatedfor species with multiple accessions, species with compound leaves areunderlined, and the asterisk indicates an especially robust specimen ofS. evolvulifolium. The topology is based on Bayesian analyses, and branchsupport values are Bayesian posterior probabilities/maximum parsimony

bootstrap.


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from S. evolvulifolium. We fully acknowledge that S. evolvuli-folium, as we recognize it herein, is paraphyletic; however, webelieve that this circumscription of the species, a species fromwhich another lineage has evolved, is biologically realisticand is more useful than erecting several new species that can-not be reliably differentiated based on morphology to main-tain purely monophyletic species. As we have circumscribedit, S. evolvulifoliumis a widespread, but easily recognized spe-

cies and S. crassinerviumand S. loxophyllumare highly differ-entiated, easily identifiable species.

Taxonomic Treatment

Solanum sect. Herpystichum Bitter, Repert. Spec. Nov. RegniVeg. 17: 331. 1921.LECTOTYPE species: S. trifoliumDunal (designated by Seithe 1962).

Herbaceous to woody, ground-trailing to climbing vines,rooting at the nodes; stems and leaves glabrous to pubes-cent, the trichomes, if present, multicellular and unbranched.Sympodial units plurifoliate or unifoliate. Leaves simpleor pinnately compound, membranous to somewhat fleshy,sand-punctate, the leaf venation palmate or pinnate, the com-

pound leaves with 3 or 5 leaflets, with interstitial leafletslacking; pseudostipules absent. Inflorescences usually extra-axillary, rarely axillary or leaf-opposed, usually opposed byadventitious roots in the ground-trailing species, usuallyunbranched, but rarely once or twice branched, ebracteate,

bearing 3116 flowers (scars), but usually fewer than 15; pedi-cels articulated at the base. Buds globose, rounded to pointedat apex (onion-shaped). Flowers apparently all perfect. Calyx

short-lobed, the lobes membranous to fleshy; corollas stel-late to pentagonal to rotate, 720 mm in diameter; anthersyellow, more or less equal, short and stout, oblong, not con-nivent, dehiscing by large apical pores, opening into introrselongitudinal slits with age, the filaments free, glabrous; ovaryglabrous to minutely papillose. Fruit a berry, yellow, bronze-

brown, purplish-black, to orangish-red when mature, occa-sionally mottled, slightly flattened and globose to conical tospindle-shaped, or strongly flattened and rhomboid to del-toid in outline, flattened perpendicular to the septum, thestrongly flattened fruits with a narrow ridge around the mar-gin. Seeds (mature) 1.52.5 1.52 mm, rounded, teardrop-shaped to reniform, flattened, ca. 0.5 mm thick, tan to lightreddish-brown, the surface minutely rugose to granular.

Key to the Species of SOLANUMSect.HERPYSTICHUM

1. Leaves simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22. Petioles mostly 3 cm long; leaf venation palmate77. Plants pubescent; corollas rotate-stellate (divided to about the middle); high elevations (>2,000 m) . . . . . . . . . . . . . . . . . . . 2. S. dalibardiforme7. Plants glabrous; corollas stellate (divided nearly to the base); low elevations (


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venation pinnate, with 57 pairs of secondary veins, theseconspicuous and prominent abaxially, densely sand-punctate;

base rounded to truncate to cordate, sometimes oblique; mar-gins entire; apex shortly acuminate; petioles (0.2)11.5 cm,glabrous or rarely pubescent adaxially, densely white sand-punctate. Internodes 1.55.5 cm. Inflorescences 13 cm longin flower to ca. 6 cm in fruit, unbranched to branched, stem-terminal to axillary to extra-axillary, with 216 flowers (scars),

the axes glabrous; peduncle 0.10.5 cm; rachis 0.12 cm; pedi-cels 412 mm in flower, 918 mm in fruit, only slightly enlargeddistally, glabrous to rarely sparsely pubescent, spaced nearlycontiguously. Calyx 2.53.5 mm long, glabrous, the mar-gins thickened, the tube 2.53 mm long, the lobes 1.52.5 1.52 mm, deltoid, acute to acuminate at tips, white to palepink; fruiting calyx somewhat accrescent, the lobes 1.52.5 1.52.5 mm. Corolla 1.52 cm in diameter, 58 mm long, stel-late, somewhat fleshy, white, the lobes 58 2.53 mm, planarat anthesis, acute to acuminate at apices, glabrous adaxi-ally, sparsely pubescent near the apex abaxially, the marginsdensely ciliate. Stamens with filaments 11.5 mm long, gla-

brous; anthers 34 1.21.5 mm. Ovary glabrous; style 46 ca. 0.3 mm, glabrous, cylindrical, sometimes deflected to one

side of flower; stigma capitate. Fruit (immature) 0.71 0.70.9 cm, ovoid to nearly globose, slightly flattened, somewhatpointed at apex, green to pale orangish to brownish at matu-rity, glabrous. Seeds 22.2 1.82 mm, flattened-reniform,tan, the surface minutely reticulate-rugulose. Figure 1FG.

Habitat and DistributionSolanum crassinervium occurswest of the Andes in southwestern Colombia and northwest-ern Ecuador in lowland and premontane rainforest habitats,including the Mache-Chindul mountain range in northwest-ern Ecuador; 150600(1,800) m in elevation (Fig. 5 ).

PhenologyFlowering apparently occurs year-round;fruiting specimens have been collected from Jan.Feb., andSept.Dec.

Conservation StatusAccording to the IUCN red list cat-

egories (IUCN 2010), S. crassinerviumis classified as B1a+biii

(endangered). This species is restricted to rainforest habi-tats in northwestern Ecuador and extreme southwesternColombia, covering an area estimated to be considerably lessthan 5,000 km2. This area is one of the more inaccessible partsof Ecuador, but increasing exploitation of this area continuesto decrease the amount of suitable habitat for S. crassinervium(C. Aulestia, Bilsa Biological Station, pers. comm.).

EtymologyThe epithet crassinerviumdescribes the promi-

nent secondary veins that are useful in helping distinguishthis species from its closest relatives.NotesSolanum crassinervium is one of the climbing spe-

cies, and can be distinguished from the other species in thesection by the somewhat fleshy texture of the stems, leaves,and flowers, its ovate to elliptical leaves with conspicuoussecondary veins that are visible in fresh and dried material,and its occasionally branched inflorescences. This is the mostrobust species of sect.Herpystichum.

Solanum crassinervium is most similar to S. evolvulifo-lium and S. loxophyllum, but differs from both species in itsrobust habit, broadly ovate leaves (vs. mostly oblong), fleshycalyx and corolla, and inflorescences that may be simple and

branched on the same individual. It can be easily differen-

tiated from S. evolvulifoliumby its much larger leaves, peti-oles, and internodes. The leaves of both S. crassinerviumandS. loxophyllumare somewhat fleshy, and both species tend todry dark to nearly black; however, S. crassinerviumhas moresecondary veins (57 vs. 34 pairs) and these are prominentabaxially, whereas those of S. loxophyllum are often obscurewithin the fleshy leaf blade (translucent in living material andflush with the leaf surface in dried material). The often stoutand branched inflorescences of S. crassinervium differ fromthose ofS. loxophyllum, which are apparently always simple,and are slender and delicate. Furthermore, inflorescences ofS. crassinerviumare typically produced in the leafy part of thestem, as compared to those of S. loxophyllum,which are typi-cally borne on older, leafless parts of the stem.

Additional Specimens ExaminedCOLOMBIA. Nario: 1,000 m,9 May 1939 (fl),A. H. G. Alston 8547(BM); Trayecto Pialapi La Planada,110N 7758W, 1,3001,700 m, 23 Jul 1988 (fr), O. de Benavides 10150(MO); Mpio. Barbacoas, Corregimiento Altaquer, Vereda El Barro, ReservaNatural Ro amb, margen derecha del Ro amb, 118N 7808W,1,325 m, 1 Dec 1993 (fl), P. Franco et al. 4707(COL, NY); Mpio. Tumaco,Resguardo de Alb, lado izquierdo del Ro Alb, 122N 7828W, 220280 m, 12 Nov 1995 (fl, fr), B. R. Ramrez et al. 8826(NY); Mpio. Barbacoas,Resguardo Indgena de Saund, 130N, 7820W, 350 m, 21 Jan 1996 (fr),B. R. Ramrez et al. 9699 (NY).

ECUADOR. Carch: Ro Blanco drainage above Chical, ca. 12 km W ofMaldonado, 1,3001,500 m, 25 Sep 1979 (fl), A. Gentry & G. Schupp 26522(MO); Cantn Tulcn, Parroquia Tobar Donoso, Reserva Indgena Aw,Centro El Baboso, 053N 7825W, 1,800 m, 1727 Aug 1992 (fr), G. Tipazet al. 1802(MO); Cantn Tulcn, Parroquia Tobar Donoso, Reserva IndgenaAw, Centro El Baboso, 053N 7825W, 1,800 m, 1727 Aug 1992 (fr),G. Tipaz et al. 1813(BM, NY, QCNE); border area between Prov. Carchi

and Esmeraldas, 20 km past Lita on road Lita-Alto Tambo, 550 m, 25 Jun1991 (fl),H. van der Werff et al. 11972(MO, NY, QCNE). Esmeraldas: BilsaBiological Station, Montaas de Mache, 20 km NW of Quinind, 3 km Wof Santa Isabel, 022N 7945W, 600 m, 26 Sep 1994 (fr),J. R. Abbott 15256(MO, SEL); San Lorenzo, Reserva tnica Aw, Parroquia Alto Tambo,Centro de la Union, Caon del Ro Mira, 052N 7826W, 250 m, 22 Mar1993 (fl), C. Aulestia & M. Aulestia 1431(MO, QCNE); Cantn Quinind,Bilsa Biological Station, Montaas de Mache, 35 km W of Quinind, 5 km Wof Santa Isabel, reserve boundary N from Station road, between the RoCube tributary and the E-bearing boundary crossing the Ro Cube, 021N7944W, 400600 m, 26 Sep 1994 (fr),M. S. Bass & N. Pitman 68(BM, NY);Bilsa Biological Station, Montaas de Mache, 35 km W of Quinind, 5 kmW of Santa Isabel, 021N 7944W, 400600 m, 6 Dec 1994 (fl), M. S. Bass &N. Pitman 289 (MO); San Jos, km 321 along railroad from Ibarra toSan Lorenzo, 1N 78W, 350 m, 5 May 1982 (fl), B. M. Boom 1374(F, NY,QCA); Cantn Quinind, Bilsa Biological Station, Montaas de Mache,

Fig. 5. Distribution of Solanum crassinervium, S. dalibardiforme, andS. dolichorhachis.


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35 km W of Quinind, 5 km W of Santa Isabel, Monkey Bone Trail, 021N7944W, 400600 m, 15 Sep 1994 (fr), J. L. Clark & B. Adnepos 55 (MO,QCNE); Cantn Quinind, Bilsa Biological Station, Mache Mountains,35 km W of Quinind, 5 km W of Santa Isabel, 021N 7944W, 400600 m,24 Jan 1995 (fl), J. L. Clark 412(BM, NY, QCNE); Cantn Quinind, BilsaBiological Station, Mache Mountains, 35 km W of Quinind, 5 km W ofSanta Isabel, 021N 7944W, 500 m, 18 Feb 1996 (fr), J. L. Clark 2121(BM,NY, QCNE, US); Cantn Quinind, Mache-Chindul Ecological Reserve,Bilsa Biological Station, Mache Mountains, 35 km W of Quinind, 021N7944W, 500 m, 110 Jan 1997 (fr), J. L. Clark 2993 (MO, US); Cantn

Quinind, Mache-Chindul Ecological Reserve, Bilsa Biological Station,Mache Mountains, 35 km W of Quinind, 021N 7944W, 500 m, 110 Jan1997 (fr),J. L. Clark et al. 3762(MO, NY, QCNE, US); 10 km W of Lita onroad to San Lorenzo, 055N 7830W, 800 m, 12 May 1991 (fl), A. Gentryet al. 69984(GOET, MO, NY); Cantn San Lorenzo, Lita to El Cristal road,finca of Dr. La Lama, 13.5 km S of Lita, 049N 7826W, 1,2201,350 m,2 Nov 1992 (fl, fr), J. L. Luteyn et al. 14744(MO, NY, QCA, US); CantnQuinind, carretera Herrera-El Pramo (Sta. Isabel), Estacin BiolgicaBilsa, 0136.7N 794240.4W, 580 m, 18 Feb-5 Mar 1995 (fr), W. Palacioset al. 13548 (MO, NY, QCNE); Cantn Quinind, carretera Herrera-ElPramo (Sta. Isabel), suroeste de la casa de la Estacin Biolgica Bilsa,0136.7N 794240.4W, 580 m, 24 Mar 1995 (fl), W. Palacios et al. 13719(MO); Cantn Quinind, Bilsa Biological Station, Montaas de Mache,35 km W of Quinind, 5 km W of Santa Isabela, Monkey Bone trail,021N 7944W, 400600 m, 11 Dec 1994 (fl, fr), N. Pitman & M. Bass 1091(MO, NY, QCNE); San Lorenzo, Territorio Aw, centro Mataje, 11144N783429W, 200 m, 17 Nov 2000 (fl), W. Ramrez et al. 12 (NY), 15 (NY);

Bilsa Biological Station, 5 km W of Sta. Isabel, 02049N 794241W, 540m, 14 Feb 2009 (fl, fr), S. Stern 400(QCNE, UT); Bilsa Biological Station,5 km W of Sta. Isabel, 02049N 794241W, 540 m, 13 Feb 2009 (fl, fr),E. J. Tepe & S. Stern 2729 (BM, MU, NY, QCA, QCNE, UT); Eloy Alfaro,Reserva Ecolgica Cotacachi-Cayapas, Parroquia Luis Vargas Torres,Ro Santiago, Estero Angostura, 049S 7845W, 250 m, 28 Oct 1993 (fr),

M. Tirado et al. 628 (MO); Eloy Alfaro, Reserva Ecolgica Cotacachi-Cayapas, Parroquia Luis Vargas Torres, Ro Santiago, Estero Angostura,049S 7845W, 250 m, 814 Dec 1993 (fr),M. Tirado et al. 775(MO).

2. Solanum dalibardiforme Bitter, Repert. Spec. Nov.Regni Veg. 11: 484. 1913.TYPE: COLOMBIA. Quindo,region froide Quindiu, Feb (fl.),J. Goudot 19(holotype:WW0001345 [scan!]; photos of holotype [F neg. 33066]:F957755!, GG00080130!, MO1691587!).

Herbaceous vine, terrestrial or climbing. Stems slender, her-

baceous, sparsely pubescent with trichomes 0.51 mm long.Sympodial units plurifoliate. Leaves simple, the blades 1.58 35 cm, slightly longer than wide, broadly ovate, chartaceousto somewhat fleshy, sparsely pubescent adaxially, glabrous tosparsely pubescent abaxially, moderately pubescent on veinsadaxially and abaxially, the trichomes on the adaxial side ofthe veins ca. 0.2 mm, the other leaf trichomes like those ofthe stems; venation palmate, with 57 primary veins, sparselysand-punctate; base deeply cordate; margins minutely revo-lute and with small, widely spaced teeth, these often hair-tipped, often obscure; apex acuminate; petioles 211 cm,sparsely pubescent, sparsely sand-punctate. Internodes 510 cm. Inflorescences 513 cm long, unbranched, extra-axillary,with 36 flowers, the axes sparsely pubescent; peduncle 2.5

7 cm, slender; rachis 0.82.5 cm; pedicels 1535 mm in flower,unknown in fruit, slender, sparsely pubescent, spaced 58 mmapart. Calyx 34.5 mm long, the tube 11.5 mm long, the lobes13 1.21.5 mm, rounded to lanceolate, acute to acuminate attips, sparsely pubescent abaxially with short, scattered hairs,more densely pubescent at the tips; fruiting calyx unknown.Corolla 1.51.8 cm in diameter, 610 mm long, rotate-stellateto pentagonal, membranous, white to light blue, the tube 49 mm, the lobes 2.35 67 mm, broadly deltoid, narrowlyacute at tips, glabrous adaxially, sparsely pubescent abaxially,the margins ciliate apically. Stamens with filaments 0.71 mm,glabrous; anthers 22.2 ca. 1 mm. Ovary glabrous; style46ca. 0.2 mm, glabrous to minutely papillose in lower half,

cylindrical to somewhat clavate; stigma capitate. Fruits 22.5 1.21.8 cm, ovoid, slightly to markedly flattened, roundedto pointed at apex, the color unknown, glabrous. Seedsunknown. Figure 2B.

Habitat and DistributionSolanum dalibardiformeis appar-ently endemic to Colombia (Depts. Cauca, Quindo, andTolima); 2,4003,500 m in elevation (Fig. 5).

PhenologyFlowering specimens have been collected in

Apr.Aug., and Nov. The single fruiting specimen seen wascollected in July.Conservation StatusAccording to the IUCN red list cate-

gories (IUCN 2010), S. dalibardiformeis classified as D2 (vulner-able). This species occupies a restricted area in the CordilleraCentral in Colombia and is only known from three locations.Furthermore, S. dalibardiformeis only known from nine collec-tions, suggesting that it is rare.

EtymologyThe epithet dalibardiforme refers to the super-ficial similarity of the leaves and habit of this species to thegenus Dalibarda(Rosaceae).

NotesSolanum dalibardiformeis one of the ground-trailingspecies, and is one of the most distinctive species of the sec-tion. It can be distinguished from the rest of the species in this

group by its simple leaves and rotate-stellate to pentagonalcorollas. It is most similar to the Ecuadorean S. limoncochaense,

but has pubescent vegetative parts, pentagonal corollas, andoccurs in high elevation habitats. Solanum trifoliumis the onlyother member of sect. Herpystichumwith considerable inter-petalar tissue, resulting in pentagonal or rotate corollas, butis easily distinguished by its 3-foliate compound leaves. Wehave seen only one scan of a fruiting specimen of this species(L. Reyes 119, COL), and the fruits appear to be elliptical andpointed apically, but without the distinctive arrowhead shapeofS. limoncochaense, S. phaseoloides, S. pentaphyllum, and S. tri-

folium. It is not possible to determine the cross sectional shapeof the fruit from the pressed specimens available, and it is notclear whether this species has flattened fruits like the rest of

the ground-trailing species.Additional Specimens ExaminedCOLOMBIA. Cauca: Purac,

Parque Nacional Purac, 15 Jun 1974 (fl, fr), L. Reyes 119(COL). Tolima:La Suiza, Cordillera Central, 2,600 m, 11 May 1932 (fl), J. Cuatrecasas3355 (MA); Quebrada Cajamarca to Mermillon, New Quindio Trail,Cordillera Central, 14 Aug 1922 (fl), E. P. Killip 9753(NY); Along Quindohighway, between Cajamarca and summit of Divide, 2,400 m, 27 Mar 1939(fl), E. P. Killip & G. Varela 34519 (COL, US); Roncesvalles, Vereda de SanMarcos, Finca el Corazn, 5 Nov 2003 (fl), J. Mora & J. Palma 743(COL);Roncesvalles, Vereda de San Marcos, Finca el Orinoco, 5 Nov 2003 (st),

J. Mora & J. Palma 924 (COL); Toche, 2,500 m, 25 May 1942 (fl), K. VonSneidern 3121 (NY, S); Toche, 2,500 m, 25 Apr 1942 (fl), K. Von Sneidern3121bis(LL, US).

3. Solanum dolichorhachisBitter, Repert. Spec. Nov. RegniVeg. 11: 490. 1913.TYPE: ECUADOR. Guayas: Balao,

in silvestris, rarium, May 1892 (fl, fr),H. Eggers 14641(holotype: MM0111203!; isotypes: AGH00077619!, B[destroyed], LL0403275!, LE [photo!], US1324515!,WU!; photo of B isotype [F neg. 2658]: GG00080131!).

Scandent shrub or liana, climbing to 8 m or more. Stemsslender, wiry-woody, glabrous to moderately pubescent withtrichomes 0.10.2 mm long, these frequently in distinct linesalong stem. Sympodial units plurifoliate. Leaves simple,the arrangement distichous, the blades 4.411 1.34.5 cm,23 times as long as wide, ovate to elliptical, chartaceousto coriaceous to somewhat fleshy, glabrous to moderatelypubescent on the midvein adaxially, glabrous abaxially; vena-tion pinnate, with ca. 5 pairs of secondary veins, the veins


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sand-punctate; base oblique, sides of the lamina 34 mmdistant on the petiole, the two sides rounded to truncate tosomewhat cordate; margins entire; apex acuminate; peti-oles 0.20.6 cm, moderately pubescent adaxially, sand-punctate. Internodes 0.72.8 cm. Inflorescences 355 cm long,unbranched, terminal to extra-axillary to nearly leaf-opposed,with 14116 flowers (scars), the axes slender, densely pubes-cent; peduncle 25 cm; rachis 249 cm; pedicels 56 mm in

flower, slender, 1314 mm in fruit, glabrous to minutely pubes-cent, spaced 1.58 mm apart. Calyx 1.52 mm long, membra-nous, sparsely pubescent, sand-punctate, the tube ca. 1 mmlong, the lobes 0.51 ca. 2 mm, broadly rounded, roundedto shortly acuminate at tips; fruiting calyx not accrescent.Corolla 0.71 cm in diameter, 35 mm long, stellate, membra-nous, white to greenish-white, the lobes 34.5 1.52 mm, lan-ceolate, acute at tips, reflexed at maturity, glabrous adaxially,pubescent along center of petal abaxially, the margins ciliate.Stamens with filaments 0.51 mm long, glabrous; anthers ca.2.51 mm. Ovary glabrous; style 34 ca. 0.3 mm, clavate,glabrous; stigma capitate. Fruits (immature) 918 46 cm,23 times as long as wide, spindle-shaped to possibly cor-date, pointed at apex, the color unknown, glabrous. Seeds

unknown. Figure 2C.Habitat and DistributionSolanum dolichorhachisoccurs in

the Pacific and Amazonian lowlands of Ecuador and in theAmazonian lowlands of Peru; 50350 m in elevation. (Fig. 5)

PhenologyFlowering specimens have been collected inApr. and Dec.; fruits have been collected in Apr.

Conservation StatusAccording to the IUCN red list cat-egories (IUCN 2010), S. dolichorhachisis classified as B1a+biii(critically endangered). This species is only known fromfive widely scattered locations, two of which are in westernEcuador, an area that continues to experience extreme hab-itat degradation (Dodson and Gentry 1991). The location atLimoncocha is in the center of an oil field, continued devel-opment of which is encroaching on all sides of the 4,600 ha

Reserva Biolgica Limoncocha (E. J. Tepe, pers. obs.). Despiteits broad distribution, only five collections of this species areknown, suggesting that it occurs at low densities, and appearsto have been rare even at the time of collection. The label onthe type specimen notes in silvestris, rarium(and in silvis raris-simum on some of the isotypes). Solanum dolichorhachis wascollected near the Laguna de Limoncocha near the Ro Napoin 1974 (B. A. Drummond III 7329, MO); however, EJT wasunable to relocate this species during a five day visit to theReserva Biolgica Limoncocha in 2009.

Montfar (2000) listed S. dolichorhachisas a critically endan-gered species endemic to Ecuador. If the Peruvian collectionis, in fact, a population of S. dolichorhachis(see below), thenthe species is more widespread than previously believed.

Nevertheless, based on the number of collections available forthis species, we believe that it is critically endangered, espe-cially in western Ecuador where the type was collected andwhich has experienced tremendous habitat destruction.

EtymologyThe epithet dolichorhachis refers to the long,slender inflorescences of this species.

NotesSolanum dolichorhachis is a climbing species recog-nizable by the long and slender inflorescences, small stellateflowers, spindle-shaped fruits, and the simple, usually largeleaves with distinctly oblique bases. This species is most simi-lar to S. pacificum, but the wiry-woody stems, densely short-pubescent inflorescences and young stems, and oblique leaf

bases set this species apart.

The label from the type collection describes the fruits as cor-date (fructu cordato). It is difficult to know what Eggers sawin three dimensions based on the single fruit on the isotypeat A, but Eggers notes suggest that the fruits may have beenflattened and, therefore, possibly somewhat like those of theground-trailing species in shape (i.e. Figs. 1BC).

The collections of this species from eastern Ecuador dif-fer somewhat from the type in that they have longer pubes-

cence, darker stems that are densely sand-punctate (vs. lighttan, non-punctate stems), inflorescences with less dense andcoarser pubescence, and non-punctate calyces. The Peruviancollection is closer to the type in leaf shape than collectionsfrom eastern Ecuador, but the plant appears to be ratherfleshy. It is possible that the two variants from the regions eastof the Andes merit recognition at the species level; however,they have no unambiguous, qualitative differences and, sincethis species has been collected few times, there is currentlyinsufficient material upon which to base further taxonomicdecisions.

Additional Specimens ExaminedECUADOR. Los Ros: NearQuevedo, Canton Vinces, Hacienda San Jos, 030S 7921W, 50 m, 28 Oct1934 (fl), Y. Mexia 6617(UC, US). Sucumbos: Limoncocha on Ro Napo,02315S 763635W, 300 m, 1 Oct 1974 (fr), B. A. Drummond III (MO).

Orellana: Parque Nacional Yasuni, Ro Tiputini, al noroeste de al conflu-encia con el Ro Tivacuno, 038 S 7630 W, 200300 m, 25 Apr 2002 (fl),G. Villa et al. 1461(BM, F).

PERU. Loreto: Maynas, Indiana, Reserva Explorama, 328S 7250W,106 m, 9 May 1990 (fl), R. Vsquez & N. Jaramillo 13680(MO, NY).

4. Solanum evolvulifolium Greenm., Bot. Gaz. 37: 211.1904.TYPE: COSTA RICA. San Jos: La Palma,1,460 m, Sep 1898 (fl, fr),A. Tonduz 12615(lectotype, heredesignated: GHGH00077489!; possible isolectotypes:CR12615, KK000449423!, KK000449424!, MM0111204!,NYNY00138984!, USUS00027570!).

Vine, sometimes shrubby, climbing tree trunks or other veg-etation. Stems slender, woody, glabrous to densely pubescent,the trichomes typically ca. 1 mm or longer, occasionally in

distinct lines along stem. Sympodial units plurifoliate. Leavessimple, the arrangement distinctly distichous, the blades 0.55 0.33 cm, 13 times as long as wide, ovate to oblong-ovate,chartaceous to coriaceous, sand-punctate, glabrous to denselypubescent on the leaf blade adaxially and abaxially, pubes-cent on midvein adaxially with hairs 0.20.5 mm, discolored,occasionally reddish below; venation pinnate, with 34 pairsof secondary veins, the veins densely sand-punctate; basetruncate to cordate, often oblique; margins entire to undu-late, revolute; apex rounded and apiculate to acute, some-times acuminate; petioles nearly absent to 0.5 cm, glabrousto densely pubescent, densely sand-punctate. Internodes 0.51.5(3.5) cm. Inflorescences 115 cm long, unbranched, nearlyleaf-opposed to extra-axillary, with 280 flowers (scars), the

axes glabrous to pubescent with simple, uniseriate, curledhairs; peduncle 0.61.5 cm; rachis 114 cm; pedicels 410 mm in flower, green to pink, 1015 mm in fruit, glabrousto densely pubescent, spaced nearly contiguously to 8 mmapart. Calyx 1.53 mm long, conical, the tube 12.5 mm long,the lobes 0.51.5 0.81.5 mm, deltate, rounded and minutelyapiculate at tips to acute, the margins somewhat thickened,glabrous to sparsely pubescent along margins, more dense attips of the lobes, pale green to pink; fruiting calyx minutelyaccrescent. Corolla 12 cm in diameter, 58 mm long, stel-late, membranous, greenish, white, pink, to bluish-purple,sometimes mottled, the lobes 511 23 mm, lanceolate,acute at apices, glabrous adaxially and abaxially, the margins


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ciliate. Stamens with filaments ca. 1 mm long, glabrous; anthers230.81.5 mm. Ovary glabrous; style 46 ca. 0.25 mm,cylindrical to somewhat clavate, minutely papillose in lower2/3; stigma truncate to somewhat capitate. Fruit 0.61.5 0.61 cm, globose to ovoid, slightly flattened, rounded toacute at apex, glabrous, red to reddish-brown when ripe.Seeds 1.52 mm in diameter, rounded, tan, the surface rugu-lose. Figure 1HI.

Habitat and DistributionSolanum evolvulifoliumoccurs inCosta Rica, Panama, Colombia, Venezuela, Ecuador, and Peruas an epiphyte on tree trunks in rain and cloud forest habitats;(200)8002,600 m in elevation (Fig. 6).

PhenologyFlowering and fruiting apparently occurs year-round.

EtymologyThe epithet evolvulifoliumrefers to the similar-ity of the leaves of this species to some species of Evolvulus(Convolvulaceae).

NotesSolanum evolvulifolium, a climbing species, is rec-ognizable by its characteristic distichous leaf arrangement(Fig. 1H) and branching pattern. The main stem of this speciesis most often encountered climbing on tree trunks, attachedwith adventitious roots at the nodes; secondary branches

extend away from the main stem. Higher order branches aretypically distichous and arise at ca. 45 angles, often giving theplant a characteristic, flattened appearance. The leaves oftendiminish in size along a branch, occasionally to a branchingpoint or inflorescence, and then increase in size again. Thepedicels, calyx and corolla are frequently pinkish-white togreenish-pink. This species can be distinguished from othersimple leaved, viny members of sect.Herpystichumby the uni-formly short internodes, somewhat coriaceous leaves (insteadof chartaceous or somewhat fleshy), and the flattened aspect ofthe branches that results from the distichous leaves branches.

South American collections are more variable than thosefrom Central America in leaf shape and, especially, the degreeof pubescence. The calyx lobes of Central American collec-

tions are broadly ovate and rounded apically, whereas thelobes of most South American collections are deltoid in shape

and acute apically. These calyx characters, however, are notsufficiently uniform, nor are they correlated with other char-acters that might justify segregation of S. evolvulifolium intoone or more additional species. Several extreme forms fromSouth America are included here, and these include especiallyrobust forms from Colombia and Ecuador and an especiallypubescent form from Ecuador. It is possible that the differ-ences merit specific status, but both forms are represented by

only a few collections and are thus included within a broadlydefined and variable, yet easily identifiable S. evolvulifolium.Furthermore, the three sequenced accessions of S. evolvulifo-lium, including one accession of the robust form and two ofthe standard form, form a monophyletic group together withS. crassinerviumand S. loxophyllum (Fig. 4). The robust formofS. evolvulifolium is strongly supported as sister to CentralAmerican accessions ofS. evolvulifolium lending support forits inclusion in a morphologically variable, yet easily recog-nizeable S. evolvulifolium.

There is some confusion about the numbering and collec-tor of the lectotype collection ofS. evolvulifolium. Greenmansprotologue states that the collector and number is Pittier 7413.However, the handwritten labels on the K specimens give the

collector as Tonduz. They do not have a collector number, butthey do have a herb. nat. Cost. number 12615 (Herbario delMuseo Nacional de Costa Rica). Nevertheless, there is suffi-cient overlap of label information on different specimens todetermine that they are all clearly part of the same collection.All labels list Tonduz name and the herb. nat. Cost. number.Furthermore, it seems that Tonduz was the actual collector,

but that Pittier distributed the specimens under his own set ofnumbers (Dauphin Lpez 2009; B. Hammel, pers. comm.). Forthese reasons, we refer to the collection as Tonduz 12615.

Greenman cited two syntypes, Tonduz 12615(as Pittier 7413)and Werckl 11599, in his original description of S. evolvuli-

folium. Tonduz 12615 is chosen as the lectotype because thiscollection has many duplicates, whereas only a single speci-

men is known of the Werckl collection. The GH specimenofTonduz 12615is chosen as the lectotype because Greenmanwas at GH in 1904 and this is likely the specimen that he usedin his description of S. evolvulifolium. Furthermore, the GHspecimen gives the altitude as 1460 m, the altitude stated inthe protologue. The rest of the Tonduz 12615collections reportan altitude of 1,542 m and therefore they are listed above aspossible isolectotypes.

Representative Specimens ExaminedCOSTA RICA. La Palma, 1,500 m,Nov 1897 (fr), K. Werckl 11599 (GH). Alajuela: Reserva BiolgicaMonteverde, 1019N 8443W, 820 m, 22 Oct 1988 (fl), E. Bello 468(F, MO);Reserva Forestal de Arenal, 1018N 8442W, 850900 m, 28 Feb 1990 (fl),E. Bello 1961 (MO); San Carlos, La Forma, Finca El Jilguero, 1025 25N844205W, 800 m, 22 Nov 1992 (fl), G. Herrera 5696(MO); 12 km NNWof San Ramn by road on way to San Lorenzo, 1 km S of Balsa, 1010N

8429W, 1,100 m, 25 Apr 1983 (fl), R. L. Liesner & E. Judziewicz 14936(MO,WIS). Cartago: Beside Ro Villegas, valley of Ro Grande de Orosi, 942N8347W, 1,620 m, 11 Jan 1970 (fl), R. W. Lent 1849(F, MO, U); MonumentoNacional Guayabo, 958N 8341W, 26 Jan 1993 (fl), G. Rivera 2054(F, K);growing in dense upland rain forest about 5 km SW of Tapanti, 947N8355W, 1,500 m, 17 Aug 1967 (fr), J. Taylor & C. Taylor 4472 (MO, NY).Heredia: Vicinity of Colonia Virgen del Socorro, 1017N 8410W, 900 m,10 Aug 1975, (fl), J. Utley & K. Utley 2824 (F); Parque Nacional BraulioCarrillo, 101550N 8405W, 1,2001,400 m, 13 Nov 1986 (fl),M. Grayum &G. Herrera 7881 (MO). Limon: Gupiles, Los Angeles, San Miguel,100420N 835040W, 1,300 m, 21 Feb 1990 (fl),A. Chacn et al. 744(MO);Parque Nacional Cordillera de Talamanca, 92230N 831410W, 1,700 m,24 Mar 1993 (fl), A. Fernndez 818 (BM, MO); El Progreso, 94720N,830730W, 1,600 m, 24 Apr 1989 (fl), G. Herrera & A. Chacn 2771(F, MO, NY); San Jose: road between Alto La Palma and Bajo La Hondura,1,400 m, 24 Feb 1978 (fl), F. Almeda & K Nakai 3913 (MO); Prez Zeledn,Fig. 6. Distribution of Solanum evolvulifolium.


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Savegre, 931N 8351W, 1,900 m, 3 Aug 1994 (fl, fr), G. Herrera et al. 7253(K);woods near Quebrada Vargus, Alto La Palma, 1004N 8359W, 1,400 m,31 Mar 1974 (fl), R. W. Lent 3852(AAU, F, MO).

PANAMA. Bocas del Toro: N slope of Cerro Horqueta, 84924N822654W, 6,0007,000 ft, 5 Aug 1947 (fr), P. H. Allen 4995(F, G, U); alongroad from Fortuna Dam, towards Chiriqu Grande, 2.2 miles N of thecontinental divide, 845N 8215W, 800 m, 12 Mar 1985 (fl), G. McPherson6815(MO, WIS). Chiriqui: Boquete, Bajo Chorro, 850N 8229W, 6,000 ft,12 Jan 1938 (fl) M. E. Davidson 112(GH, F, US). Chiriqu/Bocas del Toro:Zona Protectora Palo Seco, along continental divide, 847N 8213W,

1,1001,300 m, 11 Aug 2000 (fl), S. Knapp & J. Mallet 9178(BR, MO).COLOMBIA. Antioquia: Ro Caldera, 558N, 7458W, Jan 1953(fl), Bro. Daniel 4498 (US). Caldas: La Finca, Quindo, 427N 7540W,3,200 m, Feb 1937 (st), E. Dryander 2136 (US); Santa Cecelia, Cordilleraoccidental,vertiente occidental, 518N 7613W, 800 m, 29 Dec 1945 (fl),K. von Sneidern 5524(F). Cauca: El Tambo, Parque Nacional Munchique,23640N 745410W, 2,570 m, 20 Jul 1993 (fl, fr), G. Lozano et al. 6961(COL, MA); El Cairo, Cerro del Ingls, Cordillera Occidental, 445N7613W, 2,260 m, 5 Jan 1987 (fl), F. A. Silverstone Sopkin et al. 2967(NY); ElTambo, La Costa, 44N 771W, 1,500 m, 25 Mar 1938 (infl), K. von Sneidern1663(F, G, S, US). Choc: San Jose del Palmar, Hoya del Rio Torito, FincaLos Guaduales, 502N 7622W, 630 m, 6 Mar 1980 (fl), E. Forero et al. 6793(COL, MO); 6 km E of Ro Pato, ca. 48 km W of Las Animas on the PanAmerican Highway, 520N 7649W, 250 m, 11 Jan 1979 (fl, fr),A. H. Gentry &

A. Renteria 24016 (AAU, MO). Nario: Cordillera Occidental, Finca LaPlanada, near Chucunes, 111N 7758W, 1,950 m, 13 Jan 1981 (fr), A. H.Gentry et al. 30549(COL, MO, NY); Junin-Barbacoas road, 210 km. N of

Junin, 130N 7810W, 900 m, 26 Jul 1986, (st) , A. H. Gentry et al. 55333(MO). Putumayo: Villagarzn, Carretera a Pto. Asis, 4 May 1994 (st),J. L.Fernndez 11431(COL); Punto de Buenos Aires, Cerro Portachuelo, 022N7501W, 2,080 m, 27 Jul 1964 (fl), D. D. Soejarto 1139(GH). Risaralda: Apa,Vereda La Cumbre, 5842N 76046W, 2,285 m, 24 Feb 1983 (fl),J. H. Torres2216(COL); Pereira, La Pastora, 2,500 m, 20 Jan 1998 (fl), G. Vargas 4472(COL). Valle: Cordillera occidental, Hoya del Ro Digua, La Elsa, 315N7051W, 1,0001,200 m, 9 Nov 1943 (f l, fr),J. Cuatrecasas 15313(F, US).

VENEZUELA. Mrida: Rivas Dvila, 2227 km S of Tovar along rd. toCanagu, 814N 7145W, 2,1002,256 m, 16 Apr 1984 (fl, fr), J. L. Luteynet al. 9960(NY).

ECUADOR. Carchi: Mira, Norte del Carmen, Camino a Chical, 017N7813W, 2,000 m, 10 Feb 1992 (fl), W. A. Palacios et al. 9753 (MO); Tulcan,Reserva Indgena Aw, 053N 7825W, 1,800 m, 17 Aug 1992 (fr), G. Tipazet al. 1923(MO). Carchi/Esmeraldas: Near Lita, 052N 7827W, 600 m,19 May 1987 (fl).H. Van der Werff et al. 9493(MO). Cotopaxi: Sigchos, TriunfoGrande, 03222S 785859W, 2,349 m, 6 Aug 2003 (fl, fr),J. E. Ramos Perez

et al. 7061(NY); Pujil, Reserva Biolgica Los Ilinizas, 05845S 790653W,1,725 m, 10 Aug 2003 (st), F. A. Silverstone Sopkin et al. 10027 (MO, NY).Esmeraldas: Road Lita-Alto Tambo, ca. km 17.8, 051N 7829W, 850 m,28 Sep 1991 (fl), B. Ollgaard 99154(AAU). Loja: Cerro de Celica, Celica -Guachanam, km 2.7, 40546S 795645W, 2,250 m, 12 Apr 1994 (fl, fr), P.

M. Jorgensen et al. 93(NY). Napo: Parque Nacional Sumaco y Comunidadde Pacto Sumaco, 03856S 773549W, 1,5501,700 m, 26 Apr 1997 (fr),A.

Alvarez et al. 2017(MO); Quijos, Cosanga, Yanayacu Biological Station andCenter for Creative Studies, 03555S 775322W, 2,200 m, 18 Aug 2005(fl),J. L. Clark et al. 9438 (BM, NY, US); Km 2, carretera nueva Cotundo Coca, 042N 7742W, 1,130 m, 5 Aug 1984 (fl), C. H. Dodson et al. 15031(NY, MO). Pichincha: Reserva ecolgica Ro Guajalito, km 59 de la carreteraantigua Quito-St. Domingo de los Colorados, 01353S 744810W, 1,8002,000 m, 4 May 2000 (fl, fr), I. Tapia 1254(SEL). Sucumbios: El Reventador,colecciones en mbas mrgenes del Rio Reventador, 002N 7741W,1,850 m, 6 Oct 1990 (st),J. Jaramillo & E. Grijalva 12933(AAU, NY).

PERU. Amazonas: Monte Virgen, 50 m frente la comunidad de

Caterpiza, 355S 7742W, 200 m, 30 Aug 1979, (fl, fr), V. Huashikat 268(MO).

5. Solanum limoncochaenseTepe, J. Bot. Res. Inst. Texas 3:516. 2009.TYPE: ECUADOR. Sucumbos: Limoncocha,Reserva Biolgica Limoncocha, in wet primary forestnear NW corner of lake, 250 m, 22 Jan 2009 (fl, fr), E. J.Tepe & S. Stern 2627(holotype: QCA!; isotypes: BM!, MO!,MU272379!, NYNY01163478!, QCNE!, US!, UT!).

Herbaceous vine, terrestrial or climbing to ca. 1 m fromground. Stems slender, glabrous. Sympodial units plurifoliate.Leaves simple, the blades 3.57 3.58 cm, slightly wider thanlong, rounded, somewhat fleshy, moderately sand-punctate,glabrous; venation palmate with 5(7) primary veins, these

sparsely sand-punctate; base cordate; margins entire, slightlyrevolute on some leaves; apex rounded to obtuse to shortlyacuminate; petioles 316 cm, moderately sand-punctate, gla-

brous. Internodes 2.520 cm. Inflorescences 48 cm long,unbranched, extra-axillary, with 23 flowers, the axes gla-

brous; peduncle 1.34.5 cm, slender; rachis 0.91.5 cm; pedi-cels 1525 mm in flower, 2530 mm in fruit, slender, glabrous,spaced ca. 15 mm apart. Calyx 2.54.2 mm long, the tube

11.5 mm long, the lobes 1.52 ca. 1.2 mm, rounded, acumi-nate at tips, glabrous to sparsely pubescent abaxially, denselypubescent adaxially, purplish; fruiting calyx slightly accres-cent, the lobes 11.2 1.52 mm, truncate-acuminate. Corolla11.6 cm in diameter, 58 mm long, stellate, membranous,white, the tube 1.52 mm, the lobes 610 1.53 mm, lanceo-late, narrowly acute at tips, the apex papillose adaxially andabaxially, the margins ciliate apically. Stamens with filamentsca. 1 mm, glabrous; anthers 22.5 ca. 1 mm. Ovary sparselypapillose; style 22.5 ca. 0.3 mm, straight, cylindrical, stout,sparsely papillose in lower half; stigma capitate, somewhat2-lobed. Fruits 13 0.63.2 cm, ovoid-rhomboid, flattened,the apex truncate to emarginate, greenish-brown to purplishnear apex when immature, bronze-brown when mature,

sparsely pubescent with hairs


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did not reveal any additional populations (E. J. Tepe, pers.obs.). There is also a continuing decline in suitable habitatsin the area due to deforestation for additional oil explora-tion, and an increase in the local population resulting fromoil-associated jobs (H. Moya, Reserva Biolgica Limoncocha,pers. comm.).

EtymologySolanum limoncochaense is named for theLaguna de Limoncocha in western Ecuador, the only known

collection locality of this species.NotesSolanum limoncochaenseis one of the ground-trailingspecies and can easily be recognized by its simple, palmatelyveined leaves, glabrous vegetative parts, and stellate flowers.It is most similar to the Colombian S. dalibardiforme, whichis pubescent throughout, has rotate-stellate corollas, andappears to be a strictly high-elevation species.

This species grows in dense patches on the rainforest floorand over fallen trees. It is a weak climber, and plants in thefield were encountered climbing tree trunks up to ca. 1 mfrom the ground (E. J. Tepe and S. Stern, pers. obs.).

Additional Specimens ExaminedECUADOR. Sucumbos: Limon-cocha on Ro Napo, 300 m, 18 Oct 1974 (fl), B. A. Drummond 7350(MO);environs of Limoncocha, 240 m, 16 Jun 1978 (fl, fr), M. T. Madison et al.5327 (AAU, F, K, MO, NY, QCA, SEL); near northwest corner of lake,

Limoncocha, Sep 1969 (fl), R. N. Mowbray 699104(MO).

6. Solanum loxophyllumBitter, Repert. Spec. Nov. RegniVeg. 11: 14. 1912.TYPE: ECUADOR. Chimborazo/Guayas: In silvis tropicalibus prope Puente de Chimbo,Sep 1891 (fr), L. Sodiro 114/20(holotype: B, [destroyed];photos of holotype [F neg. 2668]: GG00080136!, NY!,NY!; isotype: QPLSSOLA0192!).

Vine, woody, climbing understory vegetation and canopytrees to 15 m or more. Stems slender, strong and wiry, theolder stems 8-shaped in cross section, reaching ca. 3 6 mmin cross section or larger, glabrous to sparsely pubescent, thetrichomes slender, 0.81.5 mm long. Sympodial units plurifo-liate. Leaves simple, the blades 2.58 0.83.5 cm, 23 times

as long as wide, ovate to elliptical, somewhat fleshy, moder-ately sand-punctate, pubescent on the midvein adaxially withsmall trichomes 0.30.5 mm long, more rarely glabrous orpubescent across the leaf surface, glabrous to pubescent onthe veins or the entire leaf surface abaxially; venation pin-nate, with 34 pairs of secondary veins, these translucent inliving material, inconspicuous and flush with the leaf surfaceon dried specimens, moderately sand-punctate; base roundedto truncate, oblique, the sides of the lamina 12 mm distanton the petiole; margins entire; apex acute; petioles 15 mm,moderately sand-punctate, pubescent adaxially. Internodes1.52.2 cm. Inflorescences 111 cm long, simple or rarelyonce branched, extra-axillary to nearly leaf opposed, with350 flowers (scars), the axes glabrous, slender and delicate;

peduncle 0.30.5 cm, slender; rachis 210 cm; pedicels 310 mm in flower, slender, 912 mm and slightly enlargedin fruit, 912 mm, somewhat thickened apically, glabrous,spaced 16 mm apart. Calyx 2.55 mm long, glabrous exceptfor the margins which are sparsely ciliate apically, the tube11.5 mm long, the lobes ca. 1.5 1.5 mm, rounded-deltoid,shortly apiculate at the tips, pale pinkish-white; fruiting calyxonly slightly accrescent, the lobes 1.52 ca. 1.5 mm. Corolla0.81 cm in diameter, 35 mm long, stellate, membrana-ceous, white, the tube ca. 1 mm, the lobes 78 22.5 mm,lanceolate, attenuate at tips, glabrous adaxially and abaxi-ally, the margins minutely ciliate apically. Stamens with fila-ments 0.81 mm, glabrous; anthers 2.53 1.21.5 mm. Ovary

glabrous; style ca. 5 0.3 mm, cylindrical, glabrous; stigmacapitate. Fruits 11.5 0.51.1 cm, ellipsoidal to conical, theapex somewhat pointed, glabrous. Seeds ca. 2 1.5 mm, ca.0.5 mm thick, flattened, teardrop-shaped, light tan in color,the surface minutely rugose. Figure 1LM.

Habitat and DistributionSolanum loxophyllum is appar-ently endemic to the Pacific lowlands and low mountains ofwestern Ecuador; 100600(850) m in elevation (Fig. 7).

PhenologyFlowering and fruiting apparently occurthroughout the year.EtymologyThe epithet loxophyllum (loxo = oblique in

Greek) makes reference to the oblique leaf bases found in thisspecies. The leaf bases ofS. dolichorhachis, however, are muchmore markedly oblique than they are in S. loxophyllum.

Conservation StatusAccording to the IUCN red list cat-egories (IUCN 2010), S. loxophyllum is classified as B1a+biii(endangered). Although this species is common in the hab-itats where it occurs, deep shade of low to midelevationrainforests in western Ecuador, less than 1,500 km2of thesehabitats remain, and they continue to be converted to agri-cultural lands (Dodson and Gentry 1991; C. Aulestia, BilsaBiological Station, pers. comm.).

Common NamesEcuador: Chinba chuba tape (Cayapa;Kvist 40522,AAU, QCA), chiro nairamo tape (Cayapa; Kvist40437, AAU, BM), quinfo aran sili (Colorado; Kvist 40691,AAU, BM).

UsesEcuador: the leaves are used for pain relief and forhealing of open wounds at joints. Leaves are applied to theaffected joint (Kvist 40437). The flowers are crushed and rubbedon the body as a perfume (Kvist 40522). The plant is usedfor a bath (bao de caliente) when feeling cold (Kvist 40691).

NotesSolanum loxophyllum is a climbing species closelyallied to S. evolvulifolium, with which it has frequently beensynonymized. It differs from S. evolvulifoliumin its thin, butdistinctly fleshy, larger leaves, less regular branching, sinu-ous rather than more or less straight branches, older stems

that are 8-shaped in cross section, a tendency toward cauli-flory, its high-climbing habit (individuals have been seen toclimb to 12 m or more; E. J. Tepe, pers. obs.), and distributionunder 850 m (S. evolvulifoliumhas occasionally been collectedas low as 250 m, but is most frequently encountered above1,000 m).

Solanum loxophyllumcan be distinguished from other speciesof sect.Herpystichumby its fleshy leaves with inconspicuousveins (translucent in fresh material), cauliflorous inflores-cences, and the distinctive older stems that are 8-shaped incross section. The slender, delicate inflorescences emergefrom the cleft between the two halves of the 8-shaped stems.Several collections from the area around Zapallo Grande andthe Colorado community, Congoma Grande, are more robust

and are more densely pubescent than collections from otherareas. Herbarium specimens of S. loxophyllum typically drydark to nearly black, a character shared with S. crassinervium.

The provenance of the type collection is rather vague,given that the town of Puente de Chimbo is no longer extant.One references states, however, that Puente de Chimbo waslocated at the end of the Chimbo valley, where the Chimboriver leaves the mountains and enters the plains, and turnsfrom a south-flowing to a west-flowing river (Wolf 1892,p. 132). This would place Puente de Chimbo near the currenttown of Bucay (Bromley and Bromley 1975), and the localityfalls well within the geographic and altitudinal range of othercollections ofS. loxophyllum.


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Based on recent fieldwork, it appears that S. loxophyllumiswidespread and common in the Pacific lowlands of Ecuador.It seems to be tolerant of intermediate levels of disturbance,and is abundant along trails and in secondary, but shady hab-itats. Flowering is apparently rather rare, and the flowers areinconspicuous and few in number. Consequently, S. loxophyl-lumis underrepresented in herbarium collections.

Additional Specimens ExaminedECUADOR. Azuay/Caar: MantaReal, Ro Patul, Sur de la Carretera La Troncal-Zhud, 233S 7920W, 450800 m, 13 Jul 1991 (fr), R. B. Foster & B. Mitsui 13542(F, QCA). El Oro: Hcda.Daucay, Limn Playa, 328S 7845W, 500 m, 22 Apr 1994 (st), X. Cornejo &C. Bonifaz 2478(NY); 11 km W of Pinas on new road to Sta. Rosa, 850 m,8 Oct 1979 (fl), C. H. Dodson et al. 9049 (MO, SEL). Esmeraldas: Mache-Chindul Ecological Reserve, Bilsa Biological Station, Mache Mountains,35 km W of Quinind, 5 km W of Sta. Isabel, 021N 7944W, 500 m, 17 Feb1996 (fl),J. L. Clark et al. 2054(BM, MO, NY); Reserva Cotacachi-Cayapas,La Aguita, 02848N 782624W, 150 m, 26 Jun 1998 (infl), X. Cornejo &C. Bonifaz 6389(NY); Ro Cayapa, Zapallo Grande, 1 km upriver Ro ZapalloGrande, 048N 7854W, 100 m, 29 Jun 1982 (infl), L. P. Kvist 40437(AAU,BM); Ro Cayapa, Zapallo Grande, 1 km upriver Ro Zapallo Grande,048N 7855W, 100 m, 3 Jul 1982 (st), L. P. Kvist 40522(AAU, QCA); envi-rons of Lita, on the Ibarra-San Lorenzo RR, 550650 m, 10 Jun 1978 (fl),M.T. Madison et al. 5154(F, QCA); Mache-Chindul Ecological Reserve, BilsaBiological Station, Mache Mountains, 35 km W of Quinind, 5 km W ofSta. Isabel, 021N 7944W, 500 m, 13 Feb 2009 (fl), E. J. Tepe & S. Stern2726(QCNE, UT); Reserva Ecolgica Cotacachi-Cayapa, Charco Vicente,Ro San Miguel, 043N 7853W, 200 m, 2031 Sep 1993 (fr),M. Tirado et al.391(MO, QCNE). Los Ros: Ro Palenque Biological Station, km 56 on theQuevedo-Santo Domingo Rd, 030S 7921W, 150220, 8 Jun 1974 (fl, fr),C. H. Dodson 5543(SEL, US); Ro Palenque Biological Station, km 56 on theQuevedo-Santo Domingo Rd, 030S 7921W, 150220 m, 29 Mar 1980 (fr),C. H. Dodson & A. Gentry 10040(MO, SEL); Ro Palenque Biological Station,km 56 on the Quevedo-Santo Domingo Rd, 030S 7921W, 150220 m,5 Feb 2009 (st), E. J. Tepe et al. 2698 (QCNE, UT). Los Ros or Pichincha:El Centinela at crest of Montaas Ila on road from Patricia Pilar to 24 deMayo, 03733S 791746W, 600 m, 6 Feb 1979 (fl), C. H. Dodson 7389(MO,SEL); El Centinela at crest of Montaas Ila on road from Patricia Pilar to24 de Mayo, 03733S 791746W, 600 m, 20 Jul 1979 (fr), C. H. Dodsonet al. 8626(MO, SEL). Pichincha: Santo Domingo bypass ca. 3 km S of SantoDomingo, 530 m, 8 Apr 1980 (fl), C. H. Dodson & A. Gentry 10359(F, MO,SEL); Reserva Forestal ENDESA, Ro Silanche, 006N 7902W, 650700 m,9 Jun 1984 (fl, fr), J. Jaramillo(AAU, MO, QCNE); in the Colorado commu-nity congoma grande at km 23 on the Santo Domingo - Puerto Limonroad, 021S 7922W, 100 m, 21 Jul 1982 (st), L. P. Kvist 40691(AAU, BM).

7. Solanum pacificumTepe, J. Bot. Res. Inst. Texas 3: 512.2009.TYPE: ECUADOR. Los Ros: Centro CientficoRo Palenque, in secondary forest, 215 m, 5 Feb 2009 (fl,fr), E. J. Tepe et al. 2696(holotype: QCNE!; isotypes: BM!,MO!, NYNY01163476!, QCA!, UT!).

Herbaceous vine, climbing secondary vegetation in gaps.Stems slender, weakly herbaceous, glabrous; fertile branchtips pendent. Sympodial units plurifoliate. Leaves simple, the

blades 1419 4.58 cm, 23 times as long as wide, lanceolateto ovate, membranaceous to thinly chartaceous, moderatelyto densely sand-punctate, glabrous adaxially and abaxially;venation pinnate, with 47 pairs of secondary veins, these

densely sand-punctate; base rounded to obtuse, more or lesssymmetrical; margins entire; apex acuminate; petioles 11.5 cm, densely sand-punctate, glabrous. Internodes 1.57 cm.Inflorescences 410 cm long, slender, unbranched, extra-axillary, with 1758 flowers (scars), the axes glabrous, slen-der; peduncle 24.5 cm; rachis ca. 6 cm; pedicels 810 mm inflower, slender, 1520 mm in fruit, enlarged apically, glabrous,spaced nearly contiguously to 12 mm apart. Calyx 11.2 mmlong, glabrous to minutely and sparsely ciliate along mar-gins, the tube 0.50.7 mm long, the lobes 0.50.6 0.81 mm,rounded, rounded to weakly acuminate at tips; fruiting calyxsomewhat accrescent, the lobes 0.60.8 ca. 1 mm. Corolla0.81 cm in diameter, ca. 5 mm long, stellate, membranous,

green to white near the margins of the petals, the lobes 45 1.22.5 mm, ovate, reflexed at maturity, acute at apices, gla-

brous adaxially and abaxially, the margins ciliate. Stamenssubequal, with filaments ca. 0.8 mm long, glabrous; anthers1.520.71.2 mm. Ovary glabrous; style 44.5 0.10.2 mm,glabrous, slightly clavate; stigma truncate. Fruit (immature)ca. 0.9 0.6 cm, ovoid, somewhat flattened, pointed at apex,green, glabrous. Seeds unknown. Figure 1JK.

Habitat and DistributionSolanum pacificum occurs inprimary and secondary rainforest habitats in the Pacific low-lands of Ecuador; 50380 m in elevation (Fig. 8).

PhenologyFlowering specimens have been collectedfrom Feb.Aug.; the type collection, collected in Feb., is theonly fruiting specimen seen. It is likely that fruiting is morefrequent than the collection record indicates.

Conservation StatusAccording to the IUCN red list cate-gories (IUCN 2010), S. pacificumis classified as B1a+3iii (criti-cally endangered) and D2 (vulnerable because of restrictedarea of occupancy). This species is restricted to lowland rain-forest habitats of western Ecuador. This habitat type has suf-fered extreme degradation, and has been reduced from anestimated 32,000 km2 to ca. 1,500 km2 (Dodson and Gentry

1991). The six known collections of S. pacificum are from asmall portion of this area, and because of the extensive habi-tat destruction, it is possible that this species survives onlywithin the 0.87 km2Centro Cientfico Ro Palenque.

EtymologySolanum pacificum is named after the Pacificlowlands of Ecuador where it is endemic, and for Mara PazMoreno, the first authors wife and frequent field compan-ion. Pax, Latin for peace, is the root of both Pacific andPaz.

NotesSolanum pacificum is a climbing species recogniz-able by its completely glabrous vegetative parts; slender,weakly herbaceous stems; small, greenish flowers; and large,thin leaves. The leaves ofS. pacificumare deep purplish-greenabove with whitish veins, and are weakly to intensely pur-

ple below. The upper surfaces of fresh leaves have a distinctlyvelvety luster.

Within sect. Herpystichum, this species is most similar toS. dolichorhachis, but differs in having leaves with symmetri-cal vs. distinctly oblique leaf bases and green, herbaceous vs.tan, woody stems. It can be distinguished from other climbingspecies by the texture, shape, and size of the leaves. Solanum

Fig. 8. Distribution of Solanum pacificumand S. pentaphyllum.


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pacificumis also similar to the sympatric S. leptorhachisBitter[sect. Geminata(G. Don) Walp.] in the size and shape of theleaves, the long, slender inflorescences, and small, greenish-white, stellate flowers; however, S. leptorhachisis an upright,woody shrub with unifoliate sympodial units on floweringstems and geminate leaves at nonflowering nodes. In con-tract, S. pacificumhas plurifoliate sympodial units and alwayshas only one leaf per node (i.e. not geminate).

Additional Specimens ExaminedECUADOR. Junction of the prov-inces Bolivar, Caar, Chimborazo, and Guayas: Foothills of the westerncordillera near the village of Bucay, 1,0001,250 ft, 815 Jun 1945 (st),W. H. Camp E-3782 (MO). Los Ros: Cantn Quevedo, Centro CientficoRo Palenque, along road between Santo Domingo de los Coloradosand Quevedo at km 47, 1.7 km S of Patricia Pilar, 035S 7921W, 220 m,9 Apr 1992 (st), T. B. Croat 73807(MO); Ro Palenque Biological Station, km56 Rd. Quevedo-Santo Domingo, 150220 m, 26 Oct 1974 (st), C. H. Dodson5663 (SEL); Ro Palenque Biological Station, km 56 Rd. Quevedo-SantoDomingo, 150220 m, 7 Aug 1975 (fl), C. H. Dodson 5933(AAU, MO, QCA,SEL); Ro Palenque Field Station, half way between Quevedo and SantoDomingo de los Colorados, 200 m, 22 Feb 1974 (fl), A. Gentry 10109(MO).

8. Solanum pentaphyllumBitter, Repert. Spec. Nov. RegniVeg. 12: 70. 1913. Bassovia pentaphylla(Bitter) Pittier, Cat. Fl.Venez. 2: 356. 1947.TYPE: COLOMBIA. Tolima: Ibagu,1845 (fr),J. Goudot s. n.(holotype: GG00104280 [scan!];

photos of holotype [Morton neg. 8546]: F1589922!, MO1781232!, NY!; isotype: GG00096093 [scan!]; photos ofisotype [F neg. 23140]: F758058!, MO1691340!, NY!).

Solanum pentaphyllum Bitter var. caraboboanumBitter, Repert.Spec. Nov. Regni Veg. 12: 70. 1913. Bassovia pentaphylla(Bitter) Pittier var.caraboboana(Bitter) Pittier, Cat. Fl. Venez.2: 356. 1947.TYPE. VENEZUELA. Carabobo: Valencia,Oct 1843 (fl), N. Funck 791(holotype: GG00104281 [scan!];possible isotypes: BRBR00000988419!, PP00578798[scan!]).

Herbaceous vine, terrestrial, trailing over ground or fallenlogs. Stems slender, herbaceous, sparsely to densely pubes-cent, the trichomes 12 mm long. Sympodial units plurifoli-ate. Leaves 5-pinnate, the blades 1.27 1.58 cm, about aslong as wide, membranaceous to chartaceous, sparsely sand-punctate, sparsely pubescent adaxially with widely scattered,short, wide-diameter trichomes 0.31.5 0.10.2 mm, withthinner trichomes occasionally present on midvein adaxially,glabrous to pubescent only on the veins abaxially, the marginsentire, the rachis sparsely to densely pubescent with hairslike those of the stem, the pubescence especially dense in theadaxial groove; lateral leaflets (the most distal pair) 0.84 0.42 cm, oblique, elliptical on lower side, obovate on upperside giving the leaflets a falcate appearance, the base oblique,acute to obtuse to slightly cordate on the proximal side, theapex obtuse to rounded to minutely acuminate, the basalpair slightly smaller than the upper pair, the upper pair morestrongly oblique, the petiolules 0.52 mm on upper pair ofleaflets, 1.56 mm on lower pair, glabrous to densely pubes-cent, especially in the adaxial groove; apical leaflet 0.84.5 0.62.4 cm, obovate, rhomboid, the base acute to cuneate, theapex obtuse to shortly acuminate, sessile to shortly petiolu-late, the petiolule 0.52.5 mm, glabrous to densely pubescent,especially along adaxial groove; petioles 1.55.5 cm, sparselysand-punctate, sparsely to densely pubescent, especially inadaxial groove. Internodes 2.59 cm. Inflorescences 37 cmlong, unbranched, extra-axillary, with 13 flowers, the axessparsely pubescent; peduncle 26 cm, slender; rachis 0.51 cm;pedicels 510 mm in flower, 1522 mm in fruit, slender,sparsely pubescent, spaced 210 mm apart. Calyx 1.52 mm

long, the tube 0.51 mm long, the lobes 0.50.8 11.2 mm,deltoid, acute to acuminate and slightly thickened at tips,sparsely pubescent with short trichomes (


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1918). He also collected in northern Venezuela from the upperOrinoco to Puerto Cabello on the Caribbean coast, but he doesnot seem to have collected in Brazil.

The holotype of S. pentaphyllumvar. caraboboanumis also atG. Possible isotypes, held at BR and P, are also labeled Funck791, but the localities of these two specimens differ from theholotype.

Additional Specimens ExaminedCOSTA RICA. Alajuela: Cantn deSan Ramn, Cordillera de Tilarn, Bajo Los Rodrguez, Ro La Esperanza,Quebrada Mirasol, Finca Araya, 101830N 84350W, 500600 m,10 Mar 1993 (fl), Bello et al. 4862 (INB); Reserva Biolgica Monteverde,Finca Villalobos, 8 km S of Volcn Arenal, 1023N 8443W, 1,0001,200 m,21 Apr 1990 (fl), W. Haber & W. Zuckowski 9863(INB, NY).

COLOMBIA. Antioquia: Frontino, Corregimiento Nutibara, ReginMur, camino hacia La Blanquita, 645N 7625W, 1,440 m, 10 Jul 1986(fl), P. Acevedo-Rodrguez et al. 1203 (NY); Frontino, Corregimiento LaBlanquita, regin de Murr, va Nutibarra-La Blanquita, 645N 7625W,1,3501,450 m, 10 Jun 1988 (fl) R. Callejas Posada et al. 6525 (MO, NY);Frontino, Region of Murr, road between Nutibara and La Blanquita,0640N 7626W, 1,460 m, 10 Feb 1989 (fl, fr), J. M. MacDougal et al. 3876(NY); Parque Nacional Natural Las Orquideas, Sector de Calles, RoCalles, 631N 7619W, 1,3001,320 m, 29 Mar 1991 (fl), J. G. Ramrez &E. Muoz 4062 (MO). Choc: Slopes of Serrana del Darin E of Unga,801N 7705W, 3001,300 m, 19 Jun 1976 (st),A. Gentry et al. 16770(MO).El Valle: Ro Dagua Valley, La Margarita, 330N 7552W, 760 m, 4 Apr 1939(fl, fr), E. P. Killip 34895(COL, F, GH, NY, US). Los Llanos: Intendencia, ElMeta, near Villavicencio, 316N 7305W, 600 m, 20 Jan 1939 (fl), O. Haught2551 (COL, F, MA, US). Meta: Trayecto desde la Vereda Aguas Claras(escuela) hasta el puente colgante del Ro Ariari, 760800 m, 25 Oct 1995(fl),J. L. Fernndez 12883(COL); Cubarral, Vda. Aguas Claras alrededoresde la escuela Aguas Claras, 34742N 735437W, 855 m, 19 Nov 1995 (fl),

M. E. Morales 590(COL).VENEZUELA. Aragua: Colonia Tovar, between Maracai and Chorona,

1024N 6717W, 5,500 ft, 1856 (fl, fr), A. Fendler 2092 (GH, GOET);between Portachuelo and Ocumare, 1034N 6923W, 20 Jan 1924 (fl),H. Pittier 11379(G, NY, US); Toma de Rancho Grande, 1027N 680W (fl),H. Pittier 15311(US); Alto de Choroni, 1029N 6736W, 1,600 m, 16 Apr1967 (fl), B. Trujillo 7632 (WIS). Barinas: Bolivar, Altimira, La GallinetaCasero El Celoso near feldspar mine, 850N 7035W, 1,5001,700 m,6 Jun 1988 (fr), L. J. Dorr et al. 5444 (MO, NY); Bolivar, near feldsparmine, between La Soledad and Santo Domingo, 851N 7135W, 1,300 m,24 Nov 1984 (fl, fr),H. Van der Werff & F. Ortega 6142(MO, NY). Carabobo:Montagne La Soledad, 2,000 ft, Dec 1840(?) (st), J. Linden 1599 (G);Hacienda de Cura, near San Joaqun, 1006N 6746W, 1,300 m, 15 Aug1918 (fl, fr),H. Pittier 8031(GH, US); Soto de la selva pluvial de Borburrata,1021N 6803W, 600 m, Feb 1942 (fl, fr), F. Tamayo 2224(G, US). DistritoCapital: Along old road between Portachuelo and Peita (Petaquire)and Carayaca, between Colonia Tovar-Junquito road and HaciendaEl Limon, 1028N 6711W, 25 May 1963 (fl), J. A. Steyermark 91446 (F,NY, US, VEN). Lara: Jimnez, Between Alto del Viento to Cerro Pando,0939N 6934W, 1,400 m, 26 Oct 1982 (fl), G. Davidse & A. C. Gonzlez21180 (NY); Iribarren, Vecinidades de la Laguna Negra, Loma de LosNaranjos, Montaa de Macanillal y Fila de San Estaban, 952N 6918W,1,3001,500 m, 24 Mar 1975 (infl), J. A. Steyermark et al. 111628 (VEN);

Jimnez, Montaa Oscura, en La Briza, 952N 6921W, 1,630 m, 8 Aug1970 (fl, fr)J. A. Steyermark et al. 103578(M, US, VEN). Trujillo: 13 km ESEof Bocono, 1 km W of Guarameca, 911N 7009W, 1,600 m, 16 Mar 1982(infl), R. L. Liesner et al. 12907(MO, NY). Yaracuy: El Amparo camino abi-erto hasta la fila, 821N 7139W, 1,300 m, 19 Mar 1973 (fl), E. Diederichs108

(MO); Cerro la Chapa, 1012N 6824W, 1,250 m, 12 Apr 1999 (fl),J. R. Grant et al. 9903335 (US); Nirgua, Serrana Santa Mara - Cerro laChapa, 1012N 6835W, 1,2001,350 m, 31 Apr 1994 (fl), W. Meier et al.3919(MO).

9. Solanum phaseoloides Pol., Linnaea 41: 585. 1877.TYPE: COSTA RICA. Alajuela: Desengao, Jun 1875(fl), H. Polakowsky 147 (lectotype, here designated:BMBM000579755!; isolectotype: B [destroyed]).

Solanum olivaeformeDonn.Sm., Bot. Gaz. (Crawfordsville) 14: 28.1889.TYPE: GUATEMALA. Alta Verapz: Pansamal,3,800 ft, May 1887 (fl), H. von Tuerckheim 1226 (lecto-type, here designated: USUS00027712!; isolectotypes:GHGH00077524!, KK000449425!, NYNY00169750!).

Herbaceous vine, terrestrial. Stems slender, glabrous orrarely pubescent with slender trichomes 0.81.5 m long.Sympodial units plurifoliate. Leaves 3-pinnate, the blades1.57.5 17 cm, about as long as wide, membranous to chart-aceous, glabrous adaxially or with widely scattered to dense,wide-diameter trichomes, 0.31.2 0.10.2 mm on the leaf

blade, with trichomes like those of the stem more abundantalong veins, glabrous abaxially, the margins entire to undulate,

minutely revolute on some leaves, the rachis glabrous; lateralleaflets 0.55 15.5 cm, elliptical to rounded, the base oblique,cuneate to cordate on the proximal side, the apex obtuse toacuminate, the petiolules 0.53 mm, glabrous; apical leaflet0.8416 cm, broadly elliptical to rhomboid, the base cune-ate, the apex obtuse to acute, apiculate, the petiolule 15 mm,glabrous; petioles 1.513 cm, sparsely sand-punctate, gla-

brous or rarely pubescent. Internodes 312 cm. Inflorescences312 cm long, unbranched, extra-axillary, with (2)3(7) flow-ers, the axes glabrous; peduncle 1.55 cm, slender; rachis16.5 cm; pedicels 1025 mm in flower and fruit, slender, gla-

brous, spaced 48 mm apart. Calyx 1.52 mm long, the tube11.5 mm long, the lobes 0.51 ca. 1.5 mm, deltoid to trun-cate, acute to acuminate at tips, glabrous; fruiting calyx slightly

accrescent, the lobes 11.2 1.52 mm, truncate-acuminate.Corolla 11.5 cm in diameter, ca. 5 mm long, stellate, white,the tube 13 mm, the lobes 67 1.23 mm, lanceolate, acuteat the tips, glabrous adaxially and abaxially, the margins ciliate.Stamens with filaments 11.5 mm, glabrous; anthers 22.5 ca. 1 mm. Ovary glabrous; style 45 ca. 0.2 mm, cylindri-cal, glabrous; stigma capitate. Fruits 24 12.5 cm, ovoid-rhomboid, flattened, pointed at apex, greenish to yellowish,often mottled, glabrous. Seeds ca. 2 1.5 mm in diameter, len-ticular, light brown, the surface smooth. Figure 1E.

Habitat and DistributionSolanum phaseoloides occurs inthe understory of wet forests and in clearings from southernMexico (Chiapas), Guatemala, Belize, Honduras, Costa Rica,to Panama. It is usually terrestrial, but occasionally grows on

fallen logs or as an epiphyte; 5002,900 m in elevation. A sin-gle collection is known from Peru (Fig. 9 ).

PhenologyFlowering and fruiting occur throughout theyear.

EtymologyThe name, phaseoloides, refers to the similarleaves and viny habit of the genus PhaseolusL. (Fabaceae).

NotesSolanum phaseoloides is a ground-trailing speciescharacterized by 3-foliate leaves and stellate flowers. It is mostsimilar to S. pentaphyllumfrom which it differs by its 3-foliatevs. 5-foliate leaves. It also resembles S. trifolium,but can bedifferentiated by its white, stellate flowers and pointed leafapices vs. blue-violet, rotate flowers and rounded leaf apices.Vegetative parts of most collections are glabrous; however,plants with pubescent stems and petiol

A Revision of Solanum Section Herpystichum-Solanaceae

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